Allocation of Colony‐Level Foraging Effort in Vespula germanica in Response to Food Resource Quantity,Quality, and Associated Olfactory Cues

In social insects, selection takes place primarily at the level of the colony. Therefore, unlike solitary insects, social species are expected to forage at rates that maximize colony fitness rather than individual fitness. Workers can increase the net benefit of foraging by responding to increased resource availability, by responding more strongly to higher‐quality resources, and by decreasing the uncertainty with which nestmates find resources. Unlike many ants and social bees, no social wasp is known to utilize a nest‐based recruitment signal to inform nestmates of food location. On the other hand, wasps do learn the odor of food brought to the nest and use this cue to locate the food source outside the nest. Here, we quantify the effects of three food‐associated variables on the allocation of foraging effort in the yellowjacket Vespula germanica. We used an experimental approach to assess whether resource quantity, quality, or associated olfactory information affect the probability that a forager will leave the nest on a foraging trip. We addressed these questions by inserting a known amount of sucrose solution directly into nests and recording foraging effort (departure rate) over the subsequent hour‐long observation period. No differences were found in foraging effort because of the presence/absence of olfactory cues, but there was strong evidence that foraging effort increased in response to resource influx and resource quality. Thus, while olfactory cues are learned in the nest, only resource quality and the cue of increased amount of food in the nest factor into a forager's decision of whether or not to depart on a foraging trip. However, as prior work has shown, once a wasp forager leaves the nest, it uses the learned olfactory cues to aid in finding resources.     

Foraging preferences of ants on a heterogeneous Brazilian sandy shore habitat

1. Ants may select their food in response to nutritional needs of the colony and forage in a way that optimises a complementary nutrition. Even though resource availability is known to affect ant colony and individual health, there is still no study that has investigated the plastic preferences of ants according to spatial resource availability in naturally heterogeneous conditions. 2. Beaches are great biomes to test spatial foraging preference because a complete absence of nectaries can be found. Dorymyrmex nigra Pergande 1896 was found inhabiting a beach in southeastern Brazil, in which nectar sources are heterogeneously distributed. This study tested whether the foraging preference to sugar baits depended on the availability of nectar sources surrounding the nests. 3. We found that more D. nigra workers foraged on sugar baits when the colonies lacked naturally occurring nectar in their vicinity compared with colonies with abundant nectar nearby. 4. These results show that the foraging preference of ants depends upon resource availability. This is the first study to use a natural mosaic of resource availability to show that resource preference of ants is plastic and varies spatially.     

How long will honey bees (<Emphasis Type="Italic">Apis mellifera</Emphasis> L.) be stimulated by scent to revisit past-profitable forage sites?

Honey bees utilise floral food sources that vary temporally in their relative and absolute quality. Via a sophisticated colony organisation, a honey bee colony allocates its foragers such that the colony focuses on the most profitable forage sites while keeping track of changes within its foraging environment. One important mechanism of the allocation of foragers is the ability of experienced foragers to revisit past-profitable forage sites after a period of temporary dearth caused by, for example, inclement weather. The scent of past-profitable forage within the colony brought back by other foragers is sufficient to reactivate these experienced foragers. Here I determine for how long bees react to the scent of a past-profitable forage site. I show that the ability of foragers to revisit the location of a past-profitable food source diminishes rapidly over a period of 10 days, until no forager reacts to the cue (scent). I discuss the implications of these findings with respect to the colony’s ability to react rapidly to changing foraging conditions.     

To be or not to be faithful: flexible fidelity to foraging trails in the leaf‐cutting ant Acromyrmex lobicornis

1. Ants using trails to forage have to select between two alternative routes at bifurcations, using two, potentially conflicting, sources of information to make their decision: individual experience to return to a previous successful foraging site (i.e. fidelity) and ant traffic. In the field, we investigated which of these two types of information individuals of the leaf‐cutting ant Acromyrmex lobicornis Emery use to decide which foraging route to take. 2. We measured the proportion of foraging ants returning to each trail of bifurcations the following day, and for 4–7 consecutive days. We then experimentally increased ant traffic on one trail of the bifurcation by adding additional food sources to examine the effect of increased ant traffic on the decision that ants make. 3. Binomial tests showed that for 62% of the trails, ant fidelity was relatively more important than ant traffic in deciding which bifurcation to follow, suggesting the importance of previous experience. 4. When information conflict was generated by experimentally increasing ant traffic along the trail with less foraging activity, most ants relied on ant traffic to decide. However, in 33% of these bifurcations, ants were still faithful to their trail. Thus, there is some degree of flexibility in the decisions that A. lobicornis make to access food resources. 5. This flexible fidelity results in individual variation in the response of workers to different levels of ant traffic, and allows the colony to simultaneously exploit both established and recently discovered food patches, aiding efficient food gathering.     

Division of labour and specification of castes in the red imported fire ant Solenopsis invicta buren

Division of labour in Solenopsis invicta follows a familiar pattern: younger, smaller ants tend toward brood care while older, larger ants tend toward foraging. However, long-term observations of marked individuals reveal that length of nursing and foraging ‘careers’ and the age of transition between these activities vary considerably between and within size groups, and are related to length of life. Experiments with entire colonies show that larger ants are more likely than smaller ants to forage for insect prey. There are two main worker castes, ‘nurses’ and ‘foragers’, whose members span a wide age-size range, and a large ‘reserve’ subcaste, heterogeneous in age, size, and behaviour: reserves may nurse, forage, store liquid food, or relay food from nurses to foragers. The proportion of ants engaged in foraging decreases with colony size because many ants in large colonies are not exposed to recruitment signals.     

How patrollers set foraging direction in harvester ants

Recruitment to food or nest sites is well known in ants; the recruiting ants lay a chemical trail that other ants follow to the target site, or they walk with other ants to the target site. Here we report that a different process determines foraging direction in the harvester ant Pogonomyrmex barbatus. Each day, the colony chooses from among up to eight distinct foraging trails; colonies use different trails on different days. Here we show that the patrollers regulate the direction taken by foragers each day by depositing Dufour's secretions onto a sector of the nest mound about 20 cm long and leading to the beginning of a foraging trail. The patrollers do not recruit foragers all the way to food sources, which may be up to 20 m away. Fewer foragers traveled along a trail if patrollers had no access to the sector of the nest mound leading to that trail. Adding Dufour's gland extract to patroller-free sectors of the nest mound rescued foraging in that direction, while poison gland extract did not. We also found that in the absence of patrollers, most foragers used the direction they had used on the previous day. Thus, the colony's 30-50 patrollers act as gatekeepers for thousands of foragers and choose a foraging direction, but they do not recruit and lead foragers all the way to a food source.     

The dynamic nature of the activities of the wood ant Formica aquilonia foraging to static food resources within a laboratory habitat

ABSTRACT. A laboratory colony of the wood ant Formica aquilonia (Yarrow) was maintained under strictly controlled feeding regimens. The number of foragers at a particular site is related to the previous provisioning of the colony and the familiarity and accessibility of the resource to foragers. Under a standard regimen a predictable number of ants forage to a known site and display a typical pattern of daily activity. This pattern is examined under stable and changing circumstances. Following food-deprivation individuals increase their foray-frequency and carry larger volumes of sugar solution per foray. On days of critical food-deficit supernumerary workers are recruited. How foragers are directed to a novel food source in unfamiliar territory is examined. Following-behaviour is not involved, neither it appears are pheromones.     

Spatial foraging patterns and colony energy status in the African honey bee,Apis mellifera scutellata

The relationship between changes in foraging patterns (inferred from waggle dance activity) and colony energy status (inferred from brood rearing activity, food storage, and colony weight) was examined for the African honey bee during a period of relative resource abundance and resource dearth. When resources were more abundant mean foraging distances (about 400 m) and foraging areas (4–5 km²) were small, and colonies recruited to 12–19 different sites per day. Colony foraging ranges and sites visited increased slightly during the dearth period, yet foraging continued to be concentrated within less than 10 km². The degree to which fluctuations in foraging patterns were correlated with colony energy status varied with the availability of floral resources. During periods of relative forage abundance, increases in foraging range and number of sites visited were significantly correlated with increases in brood rearing and colony weight. In contrast, colonies examined during periods of resource dearth exhibited no correlations between foraging areas, foraging distances, and fluctuations in brood rearing, food storage, or colony weight. Thus, during dearth periods colonies may not be able to coordinate foraging patterns with changes in colony energy status.     

Colony variation in the collective regulation of foraging by harvester ants

This study investigates variation in collective behavior in a natural population of colonies of the harvester ant, Pogonomyrmex barbatus. Harvester ant colonies regulate foraging activity to adjust to current food availability; the rate at which inactive foragers leave the nest on the next trip depends on the rate at which successful foragers return with food. This study investigates differences among colonies in foraging activity and how these differences are associated with variation among colonies in the regulation of foraging. Colonies differ in the baseline rate at which patrollers leave the nest, without stimulation from returning ants. This baseline rate predicts a colony's foraging activity, suggesting there is a colony-specific activity level that influences how quickly any ant leaves the nest. When a colony's foraging activity is high, the colony is more likely to regulate foraging. Moreover, colonies differ in the propensity to adjust the rate of outgoing foragers to the rate of forager return. Naturally occurring variation in the regulation of foraging may lead to variation in colony survival and reproductive success.     

Interactions with Combined Chemical Cues Inform Harvester Ant Foragers' Decisions to Leave the Nest in Search of Food

Social insect colonies operate without central control or any global assessment of what needs to be done by workers. Colony organization arises from the responses of individuals to local cues. Red harvester ants (Pogonomyrmex barbatus) regulate foraging using interactions between returning and outgoing foragers. The rate at which foragers return with seeds, a measure of food availability, sets the rate at which outgoing foragers leave the nest on foraging trips. We used mimics to test whether outgoing foragers inside the nest respond to the odor of food, oleic acid, the odor of the forager itself, cuticular hydrocarbons, or a combination of both with increased foraging activity. We compared foraging activity, the rate at which foragers passed a line on a trail, before and after the addition of mimics. The combination of both odors, those of food and of foragers, is required to stimulate foraging. The addition of blank mimics, mimics coated with food odor alone, or mimics coated with forager odor alone did not increase foraging activity. We compared the rates at which foragers inside the nest interacted with other ants, blank mimics, and mimics coated with a combination of food and forager odor. Foragers inside the nest interacted more with mimics coated with combined forager/seed odors than with blank mimics, and these interactions had the same effect as those with other foragers. Outgoing foragers inside the nest entrance are stimulated to leave the nest in search of food by interacting with foragers returning with seeds. By using the combined odors of forager cuticular hydrocarbons and of seeds, the colony captures precise information, on the timescale of seconds, about the current availability of food.     

Context-dependent navigation in a collectively foraging species of ant, Messor cephalotes

More than 100 years of scientific research has provided evidence for sophisticated navigational mechanisms in social insects. One key role for navigation in ants is the orientation of workers between food sources and the nest. The focus of recent work has been restricted to navigation in individually foraging ant species, yet many species do not forage entirely independently, instead relying on collectively maintained information such as persistent trail networks and/or pheromones. Harvester ants use such networks, but additionally, foragers often search individually for food either side of trails. In the absence of a trail, these ‘off-trail’ foragers must navigate independently to relocate the trail and return to the nest. To investigate the strategies used by ants on and off the main trails, we conducted field experiments with a harvester ant species, Messor cephalotes, by transferring on-trail and off-trail foragers to an experimental arena. We employed custom-built software to track and analyse ant trajectories in the arena and to quantitatively compare behaviour. Our results indicate that foragers navigate using different cues depending on whether they are travelling on or off the main trails. We argue that navigation in collectively foraging ants deserves more attention due to the potential for behavioural flexibility arising from the relative complexity of journeys between food and the nest.     

Taking a trip to the shelf: Behavioral decisions are mediated by the proximity to foraging habitats in the black‐legged kittiwake

For marine top predators like seabirds, the oceans represent a multitude of habitats regarding oceanographic conditions and food availability. Worldwide, these marine habitats are being altered by changes in climate and increased anthropogenic impact. This is causing a growing concern on how seabird populations might adapt to these changes. Understanding how seabird populations respond to fluctuating environmental conditions and to what extent behavioral flexibility can buffer variations in food availability can help predict how seabirds may cope with changes in the marine environment. Such knowledge is important to implement proper long‐term conservation measures intended to protect marine predators. We explored behavioral flexibility in choice of foraging habitat of chick‐rearing black‐legged kittiwakes Rissa tridactyla during multiple years. By comparing foraging behavior of individuals from two colonies with large differences in oceanographic conditions and distances to predictable feeding areas at the Norwegian shelf break, we investigated how foraging decisions are related to intrinsic and extrinsic factors. We found that proximity to the shelf break determined which factors drove the decision to forage there. At the colony near the shelf break, time of departure from the colony and wind speed were most important in driving the choice of habitat. At the colony farther from the shelf break, the decision to forage there was driven by adult body condition. Birds furthermore adjusted foraging behavior metrics according to time of the day, weather conditions, body condition, and the age of the chicks. The study shows that kittiwakes have high degree of flexibility in their behavioral response to a variable marine environment, which might help them buffer changes in prey distribution around the colonies. The flexibility is, however, dependent on the availability of foraging habitats near the colony.     

Capital and income breeding and the evolution of colony founding strategies in ants

Ability to store resources that will be used for reproduction represents a potential life history adaptation because storage permits feeding and reproduction to be decoupled spatially and/or temporally. The two ends of a continuum involve acquiring all resources prior to reproduction (capital breeding) or acquiring all resources during the reproductive period (income breeding). Traditional life history theory examines tradeoffs between costs and benefits of such strategies, but this theory has not been integrated into life history studies of ants, even though founding queens have the analogous strategies of fully claustral (capital breeding) and semi-claustral (income breeding). This study demonstrates that facultatively semi-claustral queens of the seed-harvester ant Pogonomyrmex desertorum exhibit phenotypic plasticity during colony founding because unfed queens produced few, small minims, whereas ad libitum fed queens produced larger, heavier minims and additional brood. Fed queens also lost less mass than unfed queens despite their producing more brood. Overall, foraging provides queens with a suite of benefits that likely offset potential negative effects of foraging risk. Life history studies across a diverse array of taxa show that capital breeding is consistently associated with low availability and/or unpredictability of food, i.e., environmental conditions that favor prepackaging of reproductive resources. Such a broad and consistent pattern suggests that similar factors favored the evolution of fully claustral (capital breeding) colony founding in ants. Overall, these data suggest that ant researchers should revise their conventional view that fully claustral colony founding evolved because it eliminated the need for queens to leave the nest to forage. Instead, colony founding strategies should be examined from the perspective of environmental variation, i.e., availability and predictability of food. I also provide a functional scenario that could explain the evolution of colony founding strategies in ants. Received 16 November 2005; revised 1 March 2006; accepted 29 March 2006.     

Nest site and weather affect the personality of harvester ant colonies

Environmental conditions and physical constraints both influence an animal's behavior. We investigate whether behavioral variation among colonies of the black harvester ant, Messor andrei, remains consistent across foraging and disturbance situations and ask whether consistent colony behavior is affected by nest site and weather. We examined variation among colonies in responsiveness to food baits and to disturbance, measured as a change in numbers of active ants, and in the speed with which colonies retrieved food and removed debris. Colonies differed consistently, across foraging and disturbance situations, in both responsiveness and speed. Increased activity in response to food was associated with a smaller decrease in response to alarm. Speed of retrieving food was correlated with speed of removing debris. In all colonies, speed was greater in dry conditions, reducing the amount of time ants spent outside the nest. While a colony occupied a certain nest site, its responsiveness was consistent in both foraging and disturbance situations, suggesting that nest structure influences colony personality.     

Patterns in foraging and nesting ecology in the neotropical ant, <Emphasis Type="Italic">Gnamptogenys moelleri</Emphasis> (Formicidae,Ponerinae)

Summary This study provides quantitative field data on the natural history and foraging behaviour of the Neotropical bromeliad-nesting ant Gnamptogenys moelleri (Ponerinae) in a sandy plain forest in Southeast Brazil. The ant nested on different bromeliad species and the nests were more frequently found in bigger bromeliads. The species used a wide array of invertebrates in its diet, hunting for live prey and scavenging the majority of the items from dead animals. The food items varied greatly in size (1 to 26 mm). Hunting was always performed by solitary workers. Retrieving was performed by solitary workers (small items), or by a group of 3 to 12 workers recruited to the food source (large items). Almost all G. moelleri foraging activity was restricted to the nest bromeliad. In the warm period more ants left the nest to forage, and foraging trips achieved greater distances compared to the cool season. Trap data revealed that overall availability of arthropod prey is higher in the summer than in the winter. The opportunism in nest site use and in foraging behaviour, the small foraging area, as well as the seasonal differences in foraging activity are discussed and compared with other tropical ants.Received 30 May 2003; revised 22 September 2003; accepted 3 October 2003.     

Fitness consequences of cooperative colony founding in the desert leaf-cutter ant Acromyrmex versicolor

The evolution of cooperative colony foundation (pleometrosis)in ants has been attributed to conversion of extra resourcesinto increased competitive ability. Most cooperative foundingspecies provide these additional resources from internal fatstores; however, in those species that forage for resources, theextent and type of individual investment in multiqueen coloniesis not well understood. We compared singly- and group-foundedlaboratory colonies of the desert leaf-cutter ant Acromyrmexversicolor to investigate how cooperation affects colony survival,foraging success, and worker production. Under laboratory conditions,single foundresses were significantly less likely to initiatea successful symbiotic fungus garden, which inevitably led to colonystarvation and death. If gardens were initiated successfully,however, there was no difference in the growth trajectoriesand foraging patterns between colony types. Cooperation in thisspecies may more likely be maintained by survival benefits thanby growth rate differences, which may be constrained in groupsby individual and colony-level costs.     

Bumblebee flight distances in relation to the forage landscape

1. Foraging range is a key aspect of the ecology of 'central place foragers'. Estimating how far bees fly under different circumstances is essential for predicting colony success, and for estimating bee-mediated gene flow between plant populations. It is likely to be strongly influenced by forage distribution, something that is hard to quantify in all but the simplest landscapes; and theories of foraging distance tend to assume a homogeneous forage distribution. 2. We quantified the distribution of bumblebee Bombus terrestris L. foragers away from experimentally positioned colonies, in an agricultural landscape, using two methods. We mass-marked foragers as they left the colony, and analysed pollen from foragers returning to the colonies. The data were set within the context of the 'forage landscape': a map of the spatial distribution of forage as determined from remote-sensed data. To our knowledge, this is the first time that empirical data on foraging distances and forage availability, at this resolution and scale, have been collected and combined for bumblebees. 3. The bees foraged at least 1.5 km from their colonies, and the proportion of foragers flying to one field declined, approximately linearly, with radial distance. In this landscape there was great variation in forage availability within 500 m of colonies but little variation beyond 1 km, regardless of colony location. 4. The scale of B. terrestris foraging was large enough to buffer against effects of forage patch and flowering crop heterogeneity, but bee species with shorter foraging ranges may experience highly variable colony success according to location.     

The influence of time of day on the foraging behavior of the honeybee, Apis mellifera

The honeybee time sense, or Zeitged?chtnis, is highly adaptive, allowing bees to synchronize their foraging behavior with the peak time of daily floral nectar rhythms. Each foraging group within the honeybee colony shows a high degree of fidelity to one species of flower. Across the day, the temporal accuracy of foraging visits to experimental feeding times varies considerably, being nearly exact for morning-trained foraging groups but becoming less so for foraging groups trained later in the day. The evidence gained in this study suggests that the diel change in accuracy exhibited by foraging groups, which persists after the removal of many potential environmental time cues, is an endogenously driven behavior pattern. Furthermore, it appears that individual bees are continuously and accurately aware of the time of day, but are programmed to forage with greater anticipation to late-day food sources. Therefore, two separate processes contributing to the honeybee time sense are implicated. The first varies with time of day and determines the amount of anticipatory activity directed toward the food source. The second process is invariant across the day and is involved with the individual forager's continuous, accurate awareness of time.     

The Regulation of Ant Colony Foraging Activity without Spatial Information

Many dynamical networks, such as the ones that produce the collective behavior of social insects, operate without any central control, instead arising from local interactions among individuals. A well-studied example is the formation of recruitment trails in ant colonies, but many ant species do not use pheromone trails. We present a model of the regulation of foraging by harvester ant (Pogonomyrmex barbatus) colonies. This species forages for scattered seeds that one ant can retrieve on its own, so there is no need for spatial information such as pheromone trails that lead ants to specific locations. Previous work shows that colony foraging activity, the rate at which ants go out to search individually for seeds, is regulated in response to current food availability throughout the colony's foraging area. Ants use the rate of brief antennal contacts inside the nest between foragers returning with food and outgoing foragers available to leave the nest on the next foraging trip. Here we present a feedback-based algorithm that captures the main features of data from field experiments in which the rate of returning foragers was manipulated. The algorithm draws on our finding that the distribution of intervals between successive ants returning to the nest is a Poisson process. We fitted the parameter that estimates the effect of each returning forager on the rate at which outgoing foragers leave the nest. We found that correlations between observed rates of returning foragers and simulated rates of outgoing foragers, using our model, were similar to those in the data. Our simple stochastic model shows how the regulation of ant colony foraging can operate without spatial information, describing a process at the level of individual ants that predicts the overall foraging activity of the colony.     

Assessing the effects of sodium on fire ant foraging in the field and colony growth in the laboratory

相似文献